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Arthropoda Selecta 12 (2): 6793

ї ARTHROPODA SELECTA, 2003

Cladocera (Crustacea: Branchiopoda) from the Pantanal (Brazil) Ветвистоусые ракообразные Cladocera (Crustacea: Branchiopoda) Пантаналя (Бразилия) Werner Hollwedel1, Alexey A. Kotov2 & Gerd-Oltmann Brandorff3 В. Холлведел1, А. А. Котов2, Г.-О. Брандорфф3
1 2 2 3

Oldenburger Str. 16A, Varel, D-26316 Germany; e-mail: whollwedel@t-online.de A.N. Severtsov Institute of Ecology and Evolution, Leninsky Prospect 33, Moscow 119071 Russia; e-mail: golokot2000@mail.ru Институт проблем экологии и эволюции РАН им. А.Н. Северцова, Ленинский просп. 33, Москва 119071 Россия. Georg-Grцning-Str. 29A, Bremen, D-28209 Germany; e-mail: Gobrandorf@aol.com

KEY WORDS: Cladocera, Anomopoda, Ctenopoda, morphology, systematics, taxonomy, fauna, distribution, Brazil, Pantanal. КЛЮЧЕВЫЕ СЛОВА: Cladocera, Anomopoda, Ctenopoda, морфология, систематика, таксономия, фауна, распределение, Бразилия, Пантаналь. ABSTRACT: In 1996 and 1999, a series of samples was collected from seven localities in the southern Pantanal (Mato Grosso do Sul, Brazil). Fifty species belonging to one family of the Ctenopoda Sars, 1865, and six families of the Anomopoda Sars, 1865 have been identified. They are typical for tropical fauna of Brazil. Previous records of Cladocera from Pantanal are summarised, and a check-list of species from there is composed, consisting of 69 taxa determined to species level and 19 taxa to genera though some of these records require confirmation. The species list of the nearby Paranб region consists of 40 species. The morphology of some ilyocryptids, macrothricids, and chydorids has been studied in detail using optical and scanning electron microscopy; these results are described and illustrated. Current taxonomic problems are discussed. РЕЗЮМЕ: В течение 1996 и 1999 гг. серия планктонных и бентосных проб была отобрана в семи точках Южного Пантаналя (штат Мато Гроссо ду Сул, Бразилия). В них было найдено пятьдесят видов из одного семейства отряда Ctenopoda Sars, 1865 и шести семейств отряда Anomopoda Sars, 1865, найденные животные типичны для тропической Бразилии. Проанализированы все предыдущие находки ветвистоусых ракообразных из Пантаналя, и составлен проверочный лист, включающий 69 таксона определенных до уровня вида и 19 до уровня рода, некоторые из этих определений нуждаются в проверке. Для ближайшего района (Парана) отмечено 40 видов. Морфология некоторых представителей семейств Ilyocryptidae Smirnov, 1976 sensu Smirnov, 1992, Macrothricidae Norman & Brady, 1867 и Chydoridae Stebbing, 1902 была исследована под отпическим и сканирующим электронным микроскопом, эти результаты проиллюстрированы в данной статье. Обсуждены некоторые современные проблемы таксономии Cladocera.
Printed in 2004.

Introduction
Pantanal (Mato Grosso do Sul, Brazil) is an immense wetland in the heart of South America with an area of about 140 000 km2. Its most important environmental factor is the fluctuating level of rivers and ground waters which results in inundations of varying duration and subsequent drying up of large areas. It is the controlling factor of the structure and function of the inundation area due to the cycling of nutrients and disposability of water. The floodplain is a very productive habitat for flora and fauna. Until now there has been only a single investigation into the effects of the annual water level fluctuations on the zooplankton [Espнndola, 1996]. In a lake connected to the Paraguay River Moina minuta dominated during high water periods, while Diaphanosoma birgei dominated during low water. Zooplankton reacts differently when the water level rises and falls with fluctuating ground water level, as is the case of the Nhecolвndia region. Here there are many lakes, some of which dry up while others are saline [Mourгo, 1989; Sakamoto et al., 1999]. Cladocerans and some copepods can survive these harsh conditions by producing resting eggs that are still viable even after passing through the digestive system of higher animals. Calheiros & de Oliveira [1999] considered only 6 papers on the limnology of the Pantanal region. The first author to mention the Pantanal was Daday [1905], in his massive volume about the freshwater microfauna of Paraguay. He examined many samples collected by Prof. J. D. Anisits mainly from Paraguay, including one from Corumba, Matto Grosso, inundation pool of the River Paraguay. From this sample he determined 17 cladoceran and two copepod species. We found 14 other papers with records of microcrustacean species: Matsumura-Tundisi [1986], Mourгo [1989], Reid & Moreno [1990], Miranda [1992], Por [1995], Lima [1996], Espнndola et al. [1996], Reid (1997), Bonecker et al.


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Werner Hollwedel, Alexey A. Kotov & Gerd-Oltmann Brandorff Aquidauana/Negro Rivers (AQUI) and Miranda River (MIRA): Miranda [1992], Por [1995], Bonecker et al. [1998], Rocha & Por [1998], Bezerra et al. [1999]. This paper examines original samples from two subregions the Paraguay River and Nhecolandia of the Pantanal, summarises previous cladoceran surveys to produce a check-list for the region, and comments on the taxonomic status of some species

Study Area
Our samples were collected in the Paraguay River (samples 13, 7) and Nhecolвndia subregions (samples 46) by colleagues of the Center of Agriculture Research in the Pantanal (EMBRAPA) (samples 12) and by one of the authors (GOB). These samples have already been studied by Koste [1999], who found 216 taxa of Rotifera. Locality 1 (collected in 1996 and 1999) is Canal do Tamengo near the city of Corumbб, the outflow of the large floodplain lake Lagoa de Cбceres on the Bolivian side of the Paraguay River. The Canal do Tamengo is not a channel with strong currents but the waters flow slowly over a broad area. It has a mixture of waters from the Lagoa de Cбceres and waters from the Paraguay River which upstream leave the river channel and return through the Canal do Tamengo. The ionic content of the water is somewhat higher than that of the Paraguay River due to the release of solutes by weathering of carbonate rocks in the upland watersheds [Hamilton et al., 1999]. Locality 2 is the Paraguay River itself just above Corumbб. Water level fluctuations in 1996 were little more than two meters with high water in June/July. The hydrochemistry of this part of the river represents a mixture of several upstream tributaries of diverse chemical composition and falls in the middle of the range of variation of the Pantanal waters [Hamilton et al., 1999]. Locality 7 is Baia do Castelo, a floodplain lake on the right (west) side of the Paraguay River about 80 km north of Corumbб. The lake is intensively colonized by submerged and floating aquatic plants and receives water from many drainage channels from the surrounding higher lands. Oliveira & Calheiros [2000] studied the phytoplankton, which is dominated by the nanoplanctonic Cryptophyceae. Chemical data are given by Calheiros & Hamilton [1998], who reported that as the water of the Paraguay River flows over previously dry land there is a stimulation of decomposition rates of and leaching from aquatic materials. The resulting decrease of dissolved O2 and increase of CO2 causes fish die. How cladocerans are affected when this happens is not known. The sampling site rompesaco lies in the extreme northern part of the lake, which is intensively populated by floating vegetation and receives water through a channel from the Bolivian side. Localities 46 are from the Nhecolвndia subregion, which Hamilton et al. [1998] called the Lake District. There are numerous lakes called baias, most of which are found at an altitude of less than 100 m [Fernandes et al., 1999]. They lie in fluvial sand deposits, which in former

Fig. 1. Subregions of the Pantanal wetland (adopted from Hamilton et al. [1996] with kind permission of the E. Schweizerbartsche Velagsbuchhandlung, Stuttgart). Рис. 1. Подразделение Пантаналя на подрайоны (по Хамильтону и др. [Hamilton et al., 1996], с любезного разрешения E. Schweizerbartsche Velagsbuchhandlung, Stuttgart).

[1998], Rocha & Por [1998], Heckman [1998a, 1998b], Santos-Silva et al. [1999], and Bezerra et al. [1999]. Four of these papers, Matsumura-Tundisi [1986], Reid & Moreno [1990], Reid [1997], Santos-Silva et al. [1999], dealt only with copepods; Miranda [1992] and Bezerra et al. [1999] determined the animals only to generic level; Daday [1905], Por [1995] and Rocha & Por [1998] presented in their lists not only Cladocera and Copepoda, but also Conchostraca and Ostracoda. The Pantanal is not a uniform area. Eberhard [2000] wrote that it is a complex of ecosystems. Adбmoli [1982] differentiated several phytogeographical areas, but for our purpose we use the subregions of Hamilton et al. [1996], which are more appropiate for freshwater animals (Fig. 1). A series of papers refers to crustaceans from the following six subregions: Cuiabб River (CUIA): Heckman [1998 a, b]; Piquiri/Sгo Lourenзo Rivers (PIQU): MatsumuraTundisi [1986], Lima [1996], Santos-Silva [1999]; Paraguay River (PARA): Daday [1905], Reid & Moreno [1990], Por [1995], Espнndola et al. [1996]; Nhecolвndia (NHEC): Mourгo [1989], Reid & Moreno [1990], Por [1995], Reid [1997];


Cladocera from Pantanal times were extensive dune fields. The crests of these ancient dunes, called cordilheiras, are covered by forests. The climate is hot with a pronounced rainy season from November to March. Evaporation is high, which results in an annual water deficit [Soriano,1999]. Here there are many permanent and temporary freshwater ponds or baias and some saline lakes or salinas [Mourгo, 1989; Sakamoto, 1999]. The most dilute, and the most saline waters of the southern Pantanal are found in the Nhecolвndia subregion [Hamilton et al., 1999]. Mourгo [1989] made an intensive study of the chemistry and the phytoplankton of two ponds and a saline lake. The salina exhibit an extremely high phytoplankton production but a very low diversity of zooplankton.

69

dehyde mixture, and dissected under a stereoscopic microscope for the study of their appendages and postabdomen. Drawings were prepared using a camera lucida attached to an Alphaphot compound microscope; optical photos were taken using Leitz-Dialux microscope with a system camera and interference optics. A few specimens ofGrimaldina brazzai and Onchobunops tuberculatus were lyophilised, mounted on an aluminium stub, coated with gold, and examined under a scanning electron microscope (JEOL-840A).

Results General comments
Fifty species belonging to the orders Ctenopoda Sars, 1865 (1 family) and Anomopoda Sars, 1865 (6 families) were identified in our samples (Tab. 1). The greatest number of species (30) was found in the Baia do Castelo rompesaco, followed by Canal do Tamengo (19), Baia 38 (19) and Paraguay River (18). The most abundant species in the baias were: Diaphanosoma brevireme, Ceriodaphnia cornuta, Moina micrura, M. reticulata, Alonella dadayi, Chydorus ventricosus, Euryalona orientalis, and Notoalona globulosa. A few specimens could not be

Material and Methods
The material from the Paraguay River was collected by towing a plankton net (58 чm) for five minutes slowly behind a boat. All the other samples were taken by a series of vertical net hauls and by washing out the roots of some floating water plants. All samples were fixed in 5 % formaldehyde solution. Specimens were isolated from the samples under a stereomicroscope, placed on slides in glycerol-formal-

Table 1. Distribution of cladoceran species in water bodies of the southern Pantanal. Highest abundance of the species in a sample: ћ (13 specimens) single, ћћ (410) few, ћћћ (1125) several, ћћћћ (26100) many, ћћћћћ (more than 100) masses. Abbreviations: eph ephippium, pa postabdomen. Таблица 1. Распространение ветвистоусых ракообразных в водоемах южного Пантаналя. Обилие видов в пробе: ћ (13 экземпляра) единично, ћћ (410) немного, ћћћ (1125) среднеe, ћћћћ (26100) много, ћћћћћ (более 100) массовый вид. Условные обозначения: eph эфиппиум, pa постабдомен.
1 2 3 4 5 6 7

Canal do Tamengo, 28.3.1996, 29.4.1998

Baia 38, 1.10.1997

Baia 69, 1.10.1997

Baia 75, 1.10.1997

River Paraguay, 212 1996

Species/ L ocalities:

ORDER C TE N OPODA Sar s, 1 86 5 Sididae Bair d, 1850

Diap ha no soma brevireme Koшнn ek, 1 98 1 ri Diap han oso ma spin ulo sum H erbst 1 975 Sar silato na serr ica ud a Sar s, 1 90 1
ORDER ANOM OPODA SA RS, 1 86 5 D aph niidae Str au s, 1 82 0
ћ

ћћћћ ћ

ћћ

ћ

ћ

Ceriod ap hnia co rnu ta Sar s, 1 88 5 Ceriod ap hnia reticula ta ( Ju rin e, 18 20) Cerio da phn ia silvestrii D aday, 1 90 2 Da ph nia gess neri Herb st, 1 96 7 Simo cep halu s acutiro stratu s ( K ing, 18 53) Simo cep ha lus latiros tris Sting elin, 1 90 6

ћћћћ ћ ћћћћ ћ

ћћћћ ћ

ep h
ћ

ћћћћ ћ

ћћ

ћћ ћћ ћ

ћћ ћ

Baia do Castelo rompesaco, 23.4.1998
ћћћћ ћ ћћ ћ ћћ ћ ћ ћ

Baia 85 Salina, 1.10.1997


70

Werner Hollwedel, Alexey A. Kotov & Gerd-Oltmann Brandorff
Table 1 (continuing). Таблица 1 (продолжение).
1 2 3 4 5 6 7

Canal do Tamengo, 28.3.1996, 29.4.1998

Baia 38, 1.10.1997

Baia 69, 1.10.1997

Baia 75, 1.10.1997

River Paraguay, 212 1996

Species/ L ocalities:

ORDER C TE N OPODA Sar s, 1 86 5 Mo inidae G ould en, 1 96 7

Mo ina micrur a Ku rz, 1 87 5 Mo ina minu ta H ansen,1 899 M oina reticula ta ( D ad ay, 19 05) Moino da ph nia macleayi (K ing,1853)
I lyo cry ptidae Smir nov, 197 6 sensu Sm irn ov, 1 99 2

ћ ћћћћ ћ ћћ ћћ

ћћћ ћ ћ ћћ ћ

Ilyo cr yp tus sa rsi Sting elin, 1913 Ilyo cr yp tus sp inifer H err ick, 1 88 2
M acr oth ricidae N or man & Br ady, 1 86 7
ћ ћћћ ћ ћ

ћ

Grimald ina b razz ai Richar d, 1882 Gu ernella raphaelis Richar d, 1882 Ma crothr ix elegan s Sar s, 1 90 1 Ma crothr ix sp inos a K in g, 1 85 3 Oncho bu nop s tub ercula tus Fr yer & Pag gi, 1 97 2
Bosm inidae Sar s, 1 86 5
ћ ћ

ћћ

ћћ ћ

ћ ћћ

B os min a ha gman ni Sting elin, 1904 Bo smina tu bicen Brehm, 1953 B osmin op sis deitersi Richar d, 1895
Chy dor idae Steb bin g, 1 90 2
ћ ћ

ћћћћ ћ ћћ ћћћћ ћ ћћ ћ ћ ћћ ћ ћ ћ ћ

Alo na brasiliensis ( Berg amin, 19 35) A lona g utta ta Sar s, 1 86 2 A lon a iherin gi Sar s, 1 90 1 A lon a mon acan tha Sar s, 1 90 1 A lona o ssia ni Sinev, 1998 A lon a rectan gu la Sar s, 1 86 2 A lon a ver rucos a Sar s, 1 90 1 A lon ella dad ayi Bir ge, 1 91 0 Camp tocercu s dad ayi Sting elin, 1913 Chyd oru s eu rynotu s Sar s, 1 90 1 Chyd orus nitidulus ( Sars, 19 01) Chyd oru s pub escen s Sar s, 1 90 1 Ch yd or us ven tricosu s D aday, 1 89 8 Ch yd oru s sp. Du nhevedia o do ntop lax Sar s, 1 90 1 Ep hemero po rus h yb ridu s ( D ad ay, 19 05)

ћћ

ћ ћ ћ ћ

sh ell, pa shell

sh ell, pa
ћћ ћ

ћ ћ ћћћ ћ

ћ ћћ

shell
ћћ ћ ћ ћ ћ ћћћ ћ

shell

ћћ

shell

shell

Baia do Castelo rompesaco, 23.4.1998
ћћ ћћћ ћ ћћ ћћ ћћ ћ ћ ћћ ћ ћ ћ ћ ћћ

Baia 85 Salina, 1.10.1997


Cladocera from Pantanal

71
Table 1 (continuing). Таблица 1 (продолжение).

1

2

3

4

5

6

7

Canal do Tamengo, 28.3.1996, 29.4.1998

Baia 38, 1.10.1997

Baia 69, 1.10.1997

Baia 75, 1.10.1997
ћ

River Paraguay, 212 1996

Species/ L ocalities:

ORDER C TE N OPODA Sar s, 1 86 5 Chy dor idae Steb bin g, 1 90 2

Ep hemero po rus trid entatu s ( Berg amin, 19 39) E urya lon a orienta lis ( D ad ay, 18 98) Kar ualona muelleri (Rich ar d, 18 97) Ku rzia po lyspin a H udec, 2 00 0 Leydig ia cf . stria ta Bir ab en, 1939 Leyd igio psis o rna ta D aday, 1 90 5 No toa lona g lob ulos a ( D ad ay, 18 98) Oxyu rella long icaud is Bir ge, 1 91 0 Oxyurella sp. Pleuro xu s sp. P seud ochyd oru s glob osu s (B air d, 18 48)
NU M BE R OF T AX A
ћ ћ ћћ ћћ ћ ћћ ћћ ћ ћ ћ ћ ћ ћ

ћ

19

18

19

2

11

9

identified as they consisted of solitary individuals or shells. In the samples from Paraguay River pelagic Daphnia, Ceriodaphnia, Moina and Bosmina were dominant. The majority of species we found (39) are typical representatives of the tropical fauna (see Figs. 2, 56 for some optical photos). A significant number (27) is Neotropical,

penetrating only the southern portion of Nearctic zone (Figs. 34, 79). About half of the species collected are new records for Pantanal (Tab. 2). To date the check-list of species from Pantanal, according to our and previously published data, consists of 68 taxa determined to species level, and a further 19 taxa to genera; however, some

Table 2. Occurrence of cladoceran species in the different subregions of the Pantanal (according to our and literature data), see comments below. Таблица 2. Встречаемость ветвистоусых ракообразных в различных районах Пантаналя (по нашим и литературным данным), см. комментарии ниже.
MIRA Bonecker et al. [1998], Miranda [1992], Rocha & Por [1998] AQUI Bonecker et al. [1998], Miranda [1992], Bezerra et al. [1999] CUIA Heckman [1998a, b], Rocha & Por [1998]
X

PARA Daday [1905], Espindola et al. [1996]

C TE N OPODA Sididae

Diaph an oso ma birgei Diap ha nos oma brevireme Diap han oso ma spin ulo sum Diaph an oso ma sp.
x x x

x x x x x

PIQU Lima [1996]

Species

NHEC Mourao [1989]

PARA This paper

NHEC This paper

x

Baia do Castelo rompesaco, 23.4.1998
ћћћ ћ ћћ ћћ ћ ћћћ ћ

Baia 85 Salina, 1.10.1997

29


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Werner Hollwedel, Alexey A. Kotov & Gerd-Oltmann Brandorff
Table 2 (continuing). Таблица 2 (продолжение).
MIRA Bonecker et al. [1998], Miranda [1992], Rocha & Por [1998] AQUI Bonecker et al. [1998], Miranda [1992], Bezerra et al. [1999] CUIA Heckman [1998a, b], Rocha & Por [1998]
X X X x x x x x x x x x x x x x x x x x x x x x x x x x x x x X X x x x X X

PARA Daday [1905], Espindola et al. [1996]

C TE N OPODA Sididae

La ton ops is fa scicula ta P seudosida sp. S arsila ton a serricau da ta Sid a cr ystallin a ( 1)
ANO M OPODA Daph niidae x x

x x

Ceriod aph nia co rnu ta Ceriod aph nia reticula ta C er ioda ph nia rig au di Cerio dap hn ia silvestrii ( 2) Cerio dap hn ia sp. Da phn ia gess neri Dap hn ia sp. Meg afenestr a aur ita ( 2) Sca pho leberis mucron ata (2) Scap ho leb eris sp. Simo cephalus acutirostratus Simo cep halu s latiros tris S imocepha lus serr ulatu s Simocep halu s sp.
M o inidae

x x

x

x x

x

x

x

x x

x x x x x

Mo ina micrur a ( 3) Moin a minu ta Mo ina reticula ta Moin a sp. Moino da phn ia macleayi Mo ino dap hn ia sp.
I lyo cry ptidae

Ilyocryp tus sa rsi Ilyo cr yp tus so rdidu s ( 2) Ilyocryp tus sp inifer I lyocryptu s sp.
Ma cr oth ricidae

Grimald ina b razz ai

PIQU Lima [1996]
X

Species

NHEC Mourao [1989]

PARA This paper

NHEC This paper


Cladocera from Pantanal

73
Table 2 (continuing). Таблица 2 (продолжение).
MIRA Bonecker et al. [1998], Miranda [1992], Rocha & Por [1998] AQUI Bonecker et al. [1998], Miranda [1992], Bezerra et al. [1999] CUIA Heckman [1998a, b], Rocha & Por [1998]
x x

PARA Daday [1905], Espindola et al. [1996]

ANO M OPODA Ma cr oth ricidae

Gu ernella ra ph aelis Ma cro thrix elegan s ( 4) Macroth rix l atcorn is (2) Ma cro thrix sp inos a Ma cro thrix sp. On chob un ops tu berculatu s
Bo sm inidae

x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x X

B osmin a ha gman ni B os min a lon giros tris ( 2) Bo smina ma cro styla ( 2) B osmina ten uiros tris ( 2) Bo smina tu bicen Bosmin a sp. B osmin ops is deitersi ( 5) Bosmin op sis sp.
C hy dor ida e

A croperu s sp. Alo na br asilien sis ( 6) Alon a costa ta A lon a den tifer a ( 6) Alo na g utta ta A lon a iherin gi ( 7) Alo na int er med ia A lona mo na ca nth a Alo na o ssiani A lona rectan gu la-ty pe A lon a ver rucos a Alon a sp. Alo nella clath ratu la Alonella da dayi (8) A lon ella sp. Ca mptocercus da da yi ( 9) Chydo rus eu rynotu s C hydo rus n itidulu s

PIQU Lima [1996]

Species

NHEC Mourao [1989]

PARA This paper

NHEC This paper


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Werner Hollwedel, Alexey A. Kotov & Gerd-Oltmann Brandorff
Table 2 (continuing). Таблица 2 (продолжение).
MIRA Bonecker et al. [1998], Miranda [1992], Rocha & Por [1998] AQUI Bonecker et al. [1998], Miranda [1992], Bezerra et al. [1999] CUIA Heckman [1998a, b], Rocha & Por [1998]
x 11

PARA Daday [1905], Espindola et al. [1996]

ANOM OPODA C hy dor idae

Chyd oru s p ub escen s C hydorus sphaer icu s Ch yd oru s ven tricosu s Ch yd oru s sp. Du nhevedia o do ntop lax Ep hemero po rus h ybridus Ep hemero po rus trid entatu s E uryalona o rienta lis Ka rua lon a karu a ( 10) K aru alon a muelleri ( 10) Kur zia po lyspin a Leptorhyn chus d entifer ( 2) Leydig ia stria ta Leyd igiop sis curviros tris Leydig iop sis cf . megalops Leyd igio psis o rna ta Leydig iop sis sp. N otoa lon a glo bulo sa ( 11) Oxyu rella lo ng icaud is Oxyurella sp. P leu roxus scop uliferu s Pleuro xu s sp. P seud ochyd oru s g lob osu s
ONYCHOPODA Poly ph emidae x x x x x x x x x x x x x

x x x x x x x x x x x x x x x x x x x x x x x x x x x

Polyph emu s s p. ( 1)
Number o f taxa 44 28 24 10 13

x 35 8

1) very dubious record; 2) record must be checked; 3) including Moina micrura ciliata Daday, 1905, which is regarded as a subspecies of M. micrura Kurz, 1875, but this must be checked; 4) including records of M. superaculeata and M. triserialis, which are apparently misidentifications of M. elegans; 5) Bosminella anisitsi Daday, 1903 is a jun. syn. of Bosminopsis deitersi Richard, 1895; 6) recently transferred to the genus Alona, previously these were determined as Alonella; 7) see text; 8) Phryxura dadayi = Alonella dadayi; 9) Apparently determinations of Camptocercus australis were dealing with C. dadayi; 10) Recently separated into genus Karualona Dumont & Silva-Briano, 2000, previously these were determined as Alona, Biapertura or Alonella; 11) Earlier it was placed into Indialona or Alonella.

PIQU Lima [1996]

Species

NHEC Mourao [1989]

PARA This paper

NHEC This paper


Cladocera from Pantanal

75

Table 3. Occurrence of cladoceran species in the Pantanal and Paranб (Brazil). See comments to species below table 2. Distribution: C cosmopolitan, N Neotropical, penetrating southern portion of Nearctic zone, H Holarctic, sometimes penetrating other zones, T tropicopolitan, U unknown. Таблица 3. Встречаемость ветвистоусых ракообразных в Пантанале и Паране (Бразилия), см. также комментарии к Таб. 2. Распространение: C космополитическое, N неотропическое с проникновением в южную часть неарктической зоны, H голарктическое, изредка с проникновением в другие зоны, T циркумтропическое, U неясно.
Species CTE NOPODA Sididae Dis trib utio n Pantanal (this p aper) Pantanal (previou s p aper s, in to tal) Par ana [ L an sac-To ha et al., 19 97]

Diaph an oso ma birgei Diap ha nos oma brevireme Diaph an oso ma fl uviatile Diap han oso ma spin ulo sum Diaph an oso ma sp. La ton ops is fa scicula ta P seudosida sp. Sar silato na serr ica ud a Sid a cr ystallin a
ANOM OPODA Daph niidae

N N N N N N H x x x

x x

x x x x

x x x x x

Ceriod aph nia co rnu ta C eriod aph nia reticula ta C er iodaphnia rig au di Cerio dap hn ia silvestrii Ceriod ap hn ia sp. Da phn ia gess neri Dap hn ia sp. Meg afenestra aur ita Scap holeber is mu crona ta Scap ho leb eris sp. Simo cephalus acutirostratus Simo cep halu s latiros tris S imocepha lus serr ulatu s Simoceph alus vetulu s Simocep halu s sp.
M o inidae

C C T N N P P T N C C

x x

x x

x

x x x x x x x x x x x x x x x

Mo ina micrur a Moin a minu ta Mo ina reticula ta Moin a sp. M oino da phn ia macleayi Mo inod ap hn ia sp.
I lyo cr y ptidae

C N N T

x x x

x x x x

x

x x

Ilyocryp tus sa rsi Ilyo cr yp tus so rdidu s

N C

x x


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Werner Hollwedel, Alexey A. Kotov & Gerd-Oltmann Brandorff
Table 3 (continuing). Таблица 3 (продолжение).

Species ANOM OPODA I lyo cr y ptidae

Dis trib utio n

Pantanal (this p aper)

Pantanal (previou s p aper s, in to tal)

Par ana [ L an sac-To ha et al., 19 97]

Ilyocryp tus sp inifer I lyocryptu s sp.
Macr oth ricidae

T

x

x x

x

Grimald ina b razz ai Gu ernella ra ph aelis Ma cro thrix elegan s Macroth rix la ticorn is Ma cro thrix sp inos a Ma cro thrix sp. On chob un ops tu berculatu s
Bo sm inidae

T T N C T N N C N N N C

x x x x x x x x x x x x x x x x x x x x x

x x x x x

B osmin a ha gman ni B os min a lon giros tris Bo smina ma cro styla B osmina ten uiros tris Bo smina tu bicen Bosmin a sp. B osmin ops is deitersi Bosmin op sis sp.
C hy dor idae

x x

A croperu s sp. A cro perus h arp ae A lon a affin is Alo na br asilien sis Alon a costa ta A lon a den tifer a Alon a cf . glabr a Alo na g utta ta A lon a iherin gi Alo na inter med ia A lona monacanth a Alona ossia ni A lona rectan gu la-ty pe A lon a ver rucos a Alon a sp. Alo nella clath ratu la A lon ella dad ayi A lon ella sp. Camp tocercu s dad ayi
N x T N x C C N C N U C N C U N C T x x x x x x x

x x x x x x x x x x

x x x x x

x x x


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77
Table 3 (continuing). Таблица 3 (продолжение).

Species ANO M OPODA C hy dor ida e

Dis trib utio n

Pantanal (this p aper)

Pantanal (previou s p aper s, in to tal)

Par ana [ L an sac-To ha et al., 19 97]

Chydorus eurynotu s C hydo rus n itidulu s Chyd oru s p ub escen s C hydorus sphaer icu s Ch yd oru s ven tricosu s Ch yd oru s sp. Du nhevedia o do ntop lax Ep hemero po rus h yb ridu s Ep hemero po rus trid entatu s E urya lona o rienta lis Gra pto leb eris tes tud inar ia Ka rua lon a karu a K aru alon a muelleri Kur zia latis sima Kur zia po lyspin a Leptorhyn chus d entifer Leydig ia stria ta Leydig ia sp. Leydigio psis b reviros tris L eyd igiop sis curviros tris Leydig iop sis cf . megalops Leyd igio psis o rna ta Leydig iop sis sp. Nicsmirno vius eximiu s No toalo na g lob ulos a Oxyurella cf . cilia ta Oxyu rella lo ng icaud is Oxyurella sp. P leu roxus scop uliferu s Pleuro xu s sp. P seud ochyd oru s g lob osu s
ONYCH OPODA Poly ph emidae

T N T C T N N N T C T N C N N N N N N N T T N N N

x x x x x x x x x x x x x x x x

x x

x

x x x x

x x x x x x x x x x x x x x x x x x x x x x x x

C

Polyph emu s sp.
Num ber o f taxa

H 50

x 63 40

records are dubious while others must be confirmed by further investigations. Among the 40 species from the Paranб [Lansac-Tфha et al., 1997] there are 7 species that have not been found in the Pantanal (Tab. 3). We are sure

that some of them, for example Niscmirnovius eximius, Graptoleberis testudinaria and Oxyurella ciliata, will eventually be found there, whereasAlona affinis is, most probably, a misidentification of A. ossiani.


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Comments on selected species
A. Family ILYOCRYPTIDAE Smirnov, 1976 emend. Smirnov, 1992 1. Ilyocryptus sarsi Stingelin, 1913
COMMENTS: This species, which has recently been redescribed by Kotov et al. [2002: 208219, Figs. 160] from material from the Pantanal (Baia 75), is one of the two most common Ilyocryptus species recorded from Brazil. It is unlike the European I. sordidus (Liйvin, 1848) and most of the Brazilian records of I. sordidus (see Table 2 and 3) seem to be dealing with I. sarsi (also see Kotov et al. [2002]). Daday [1905] recorded I. sordidus in his Paraguayan samples but this is in fact I. paranaensis Paggi, 1989 (P. tifter, personal communication based on study of Dadays original material). We did not find I. paranaensis in our material.

2. Ilyocryptus spinifer Herrick, 1882
COMMENTS: Kotov & Williams [2000: 6879, Figs. 1 91] selected a neotype and redescribed I. spinifer s. str. from the North American continent. Kotov & Dumont [2000: 88 100, Figs. 1149] analyzed many populations from different countries (see intensive synonymy in these publications) including material from Pantanal (Baia 38 and Baia do Castelo). This is one of the most common anomopods in tropical water bodies of different types. It is well-differentiated from other species.

B. Family MACROTHRICIDAE Norman & Brady, 1867 emend. Smirnov, 1992 3. Grimaldina brazzai Richard, 1892 Figs. 1040.
Sars, 1901; 2831, Pl. 5; Daday, 1905: 192193; Martнnez de Ferrato, 1966: 403, Pl. 2: Fig. 5; Fryer, 1974: 236238, Figs. 128 129; Smirnov, 1976: 155156, Figs. 137140; Brandorff et al., 1982: 103, Fig. 79; Smirnov, 1992: 107109, Figs. 461468; Paggi, 1995: Figs. 8788; Elmoor-Loureiro, 1998: 24; Silva-Briano, 1998: 149151, Figs. 110.

large, significantly compressed laterally, with longitudinal keel along entire preanal margin. Preanal margin with fine spines, subdivided into two lobes, a bunch of fine setules near the boundary between the lobes; postanal and anal margins with series of plumose setae, a solitary seta at posterior edge of anus. Postabdominal setae longer than postabdomen, with distal segment half the length of the basal segment. Postabdominal claw relatively robust, with a single basal spine, two rows of setules along dorsal margin and a series of fine setules along ventral margin. Antenna I long, thin, rod-like in terms of Smirnov [1992], with a sensory seta arising at distance ? of its length from base, setules along its posterior margin, a row of robust denticles around its distal end, and 9 aesthetascs of unequal size. Antenna II long, two sensory setae of similar size at coxal part, basal segment cylindrical, all branch segments elongated, with concentric rows of small denticles. Antennal formula: setae 0-0-1-3/1-1-3, spines 0-1-0/0-0-1. Swimming setae not too long (shorter than basal segment plus rami), the largest seta (basal lateral swimming seta of endopod) asymmetrically armed with short setules distally, all other lateral and apical setae with bilaterally setulated basal segments, and asymmetrically armed distal segments, chitinous insertions within each distal segment near joint with basal segment. All spines thin, about three times shorter than segments from which they arise. Limb I with ODL bearing a single, large seta, IDL with three setae, and gnathobase I with two setae of unequal size. Other limbs as described by SilvaBriano [1998]. Size 590670 чm in our material, up to 820 чm according to Sars [1901]. Ephippial female and male: See Sars [1901]. COMMENTS: This taxon seems to be tropicopolitan, but must be revised. The most detailed revision of African and South American populations by Silva-Briano [1998] did not reveal any significant differences in their morphology, but only a very limited number of parthenogenetic females was studied.

4. Macrothrix elegans Sars, 1901
COMMENTS: This species, described from Brazil, was for a long time confused with Afro-Asiatic M. triserialis Brady, 1886. Dumont et al. [2002] pointed out intercontinental differences of members from the triserialis-group, but decided that M. superaculeata Smirnov, 1982 is a junior synonym of M. elegans. More recently Kotov et al. [2003a] studied many Neotropical populations of M. elegans (see intensive synonymy here), including material from Baia do Castelo, and concluded that M. superaculeata is a valid, but relatively rare species. We did not find it in our Pantanal samples, but it will probably be found in the future. All previous determinations of M. superaculeata must be reexamined, because Smirnovs [1992] key, which determines both species by the presence or absence of large additional spines on segments of the 4-segmented antennal branch, is not correct, as both species have these spines.

Type locality: Mayoumba et Caca Mueca, Congo, Africa Parthenogenetic female: Body subquadrangular to ovoid, relatively high and significantly compressed laterally. Border between head and valves as a very shallow depression, dorsal margin of valves straight or even concave, postero-dorsal angle distinct. Dorsal margin finely serrated; this serration is connected with specific reticulations and looks like fish scales. On lateral surface of head and valves reticulation appears as a system of dorso-ventral striae. Head triangularrounded, with short rostrum; ventral margin generally straight, with a small outgrowth like a transversal plate posterior to antenna I, a minute frontal head pore near distal extremity of rostrum, dorsal head pore completely absent. Labrum in general subquadrangular, with high, setulated projection at antero-ventral angle, naked projection at postero-ventral angle, and large, setulated distal labral plate. Compound eye small, ocellus somewhat smaller than eye, located near rostrum. Valves large, with numerous marginal setae; in anterior half of ventral margin these setae are thicker and shorter, while in posterior half longer and thinner. Postero-ventral region with series of short, setulated spines, a solitary seta located submarginally on inner surface dorsally to the series of spines. Posterior margin with fine setules. Postabdomen

5. Macrothrix spinosa King, 1853 Figs. 4155.
Richard, 1897: 286287 (laticornis); Sars, 1901: 3637; Pl. 6: Figs. 1012 (squamosa); Daday, 1905: 194195 (laticornis); Harding, 1955: 339, Figs. 4044 (laticornis); Smirnov, 1976: 7279, Figs. 4047; Smirnov, 1992: 2935, Figs. 6093; Elmoor-Loureiro, 1998: 25; Silva-Briano, 1998: 346348, Figs. 145.

Type locality: A pond between Sydney and Liverpool, now destroyed by urbanization [Smirnov, 1992], New South Wales, Australia.


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79

Figs. 29. Selected anomopods from Pantanal, Mato Grosso do Sul, Brazil: 2 Guernella raphaelis from Canal do Tamengo; 3 Onchobunops tuberculatus from Baia do Castelo; 4 Alona ossiani from Baia 38 in Nhecolвndia; 5 Chydorus ventricosus from Baia do Castelo; 6 its postabdomen; 7 postabdomen of Oxyurella longicaudis from Canal do Tamengo; 8 Karualona muelleri from Canal do Tamengo; 9 postabdomen of Alona ossiani from Baia 38. Scales: 100 чm. Рис. 29. Некоторые представители Anomopoda из Пантаналя, Мато Гроссо ду Сул, Бразилия. 2 Guernella raphaelis из Canal do Tamengo; 3 Onchobunops tuberculatus из Baia do Castelo; 4 Alona ossiani из Baia 38; 5 Chydorus ventricosus из Baia do Castelo; 6 его постабдомен; 7 постабдомен Oxyurella longicaudis из Canal do Tamengo; 8 Karualona muelleri из Canal do Tamengo; 9 постабдомен Alona ossiani из Baia 38. Масштаб: 100 чm.

Parthenogenetic female: Body ovoid in lateral view, somewhat compressed laterally, with well-developed dorsal keel. Dorsal margin finely serrated along entire length, regularly curved from tip of rostrum to rounded postero-dorsal angle. Reticulation well-expressed on head, valves, and postabdomen. Head relatively large, with relatively small, ovoid dorsal

head pore; in lateral view head triangular-round, with short rostrum, minute frontal head pore near its tip, ventral margin of head concave. Eye large, ocellus small, located closer to tip of rostrum than eye. Labrum subquadrangular, with a small projection at postero-ventral angle and relatively large distal labral plate. Valves ovoid, with serrated free margin due to


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Werner Hollwedel, Alexey A. Kotov & Gerd-Oltmann Brandorff

Figs. 1023. Grimaldina brazzai, parthenogenetic female from Baia do Castelo: 1011 adult in lateral view and its head; 12 13 setae at anterior portion and middle of ventral margin; 14 postero-ventral portion of valve; 1517 postabdomen, its preanal margin and distal portion; 1819 antenna I and its medial portion; 20 antenna II; 2122 its apical and largest (basal) lateral swimming seta; 23 thoracic limb I. Scales: 100 чm. Рис. 1023. Grimaldina brazzai, партеногенетическая самка из Baia do Castelo: 1011 вид сбоку и голова; 1213 щетинки в передней части и середине брюшного края; 14 задне-брюшная часть створки; 1517 постабдомен, его преанальный край и дистальная часть; 1819 антенна I и ее средняя часть; 20 антенна II; 2122 ее апикальная и наибольшая латеральная плавательная щетинка; 23 нога I. Масштаб: 100 чm.


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81

Figs. 2432. Grimaldina brazzai, parthenogenetic female from Baia do Castelo: 2425 adult in lateral and latero-ventral view; 2627 head in latero-ventral and dorsal view; 28 reticulation of valves; 29 armature of posterior margin of valve; 3032 postabdomen in lateral and dorsal view, and its distal portion in distal view. Scales: 100 чm for 2427, 3031; 10 чm for 2829, 32. Рис. 2432. Grimaldina brazzai, партеногенетическая самка из Baia do Castelo: 2425 вид сбоку и латеро-вентрально; 2627 голова латеро-вентрально и со спины; 28 скульптура створки; 29 вооружение заднего края створки; 3032 постабдомен, вид сбоку и со спины, и его дистальная часть. Масштаб: 100 чm для 2427, 3031; 10 чm для 2829, 32.


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Figs. 3340. Grimaldina brazzai, parthenogenetic female from Baia do Castelo: 3335 distal portion of postabdomen in dorsal view, and its setae; 3637 postabdominal claws in lateral and ventral view; 3839 antenna II and its distal segment; 40 distal portion of antenna I. Scales: 10 чm. Рис. 3340. Grimaldina brazzai, партеногенетическая самка из Baia do Castelo: 3335 дистальная часть постабдомена, вид со спины, и щетинки на ней; 3637 постабдоминальные когти, вид сбоку и с брюшной стороны; 3839 антенна II и ее дистальный членик; 40 дистальная часть антенны I. Масштаб: 10 чm.

presence of marginal triangular teeth here and marginal setae articulated exactly to these teeth. Postabdomen sub-quadrangular, not bilobed, truncated distally, without a heel basally. Transverse rows of setules on preanal margin, postanal margin very small. Postabdominal setae short, as long as postabdomen or shorter, with very short distal segment, armed with a bunch of long setules. Postabdominal claws small, armed with rows of denticles along dorsal and ventral margins. Antenna I relatively short and robust for the genus, significantly widened distally, with a short sensory seta arising at distance of ? of antenna length from its base, with robust setules distally, row of setules around distal end, nine aesthetascs of significantly different sizes. Antenna II relatively large, with two sensory setae on coxal part, robust basal segment and elongated branches with rows of denticles. Antennal formula: setae 0-0-1-3/1-1-3, spines 0-1-0-1/0-0-1. Largest (basal lateral) seta significantly longer than the rest,

unilaterally setulated, other setae armed in different manner, as is shown in Figs 5254. Limb I with ODL supplied with a large and a rudimentary seta, IDL with 3 setae of different size and series of setules, a very short seta on gnathobase I. Size 320-450 чm in our material, up to 540 чm according to Smirnov [1992]. Ephippial female: was not described from South America. Male: see Smirnov [1992]. COMMENTS. Most probably, the M. laticornis reported by Richard [1897], Daday [1905] and Harding [1955] is M. spinosa, a very common species from South America. The difference between serrated and squamose dorsal outline of the valve the main distinctive trait in all old and recently used keys is quite subjective, and is not supported by any morphometric data for these conditions. This confusion accounts for many of the misidentifications from different regions of the world. Real differences between laticornis and


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83

Figs. 4155. Macrothrix spinosa, parthenogenetic female from Baia do Castelo: 4142 adult in lateral and anterior view; 43 head; 4446 setae at antero-ventral, middle, and postero-ventral part of valve; 47 postabdomen; 48 antenna I; 49 antenna II; 5051 largest basal (lateral) seta in two specimens; 5254 swimming setae of three types in antenna II; 55 thoracic limb I. Scales: 100 чm. Рис. 4155. Macrothrix spinosa, партеногенетическая самка из Baia do Castelo: 4142 вид сбоку и спереди; 43 голова; 4446 щетики в передне-брюшной, средней и задне-брюшной части створки; 47 постабдомен; 48 антенна I; 49 антенна II; 5051 наибольшая латеральная щетинка двух самок; 5254 плавательные щетинки трех типов на антенне II; 55 нога I. Масштаб: 100 чm.


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Figs. 5661. Onchobunops tuberculatus, parthenogenetic female from Baia do Castelo: 56 adult in lateral view; 5758 head in dorsal and latero-ventral view; 59 anterior head portion with frontal head pore and supra-ocular dome; 60 dorsal head pore; 61 sculpture of valves. Scales: 100 чm for 5657, 10 чm for 5861. Рис. 5661. Onchobunops tuberculatus, партеногенетическая самка из Baia do Castelo: 56 вид сбоку; 5758 голова сбоку и латеро-вентрально; 59 передняя часть головы с фронтальной головной порой и глазным куполом; 60 спинная головная пора; 61 скульптура створок. Масштаб: 100 чm для 5657, 10 чm для 5861.

spinosa groups concern the thoracic limbs [Silva-Briano, 1998; Silva-Briano et al., 1999]. Sars [1901] discussed differences between his M. squamosa and M. laticornis, but said nothing about M. spinosa, perhaps because he did not know Kings species. Only a new, more detailed revision can reveal the true status of South American spinosa-like populations. Smirnov [1992] discussed the possibility of M. goeldi Richard, 1897 as a junior synonym of M. spinosa, but populations from the non-tropical regions of the southern South American continent (M. goeldi) must be checked.

1992: 106107, Figs. 454458; Paggi, 1995: Figs. 7778; SilvaBriano, 1998: 136138, Figs. 3140; Silva-Briano & Dumont, 2001: 2527, Figs. 3140 (Bunops); Elнas-Gutiйrrez et al., 2001: 46, Fig. 13. ? Daday, 1905: 195, pl. 12: Figs. 1517 (Macrothrix gibbera).

6. Onchobunops tuberculatus Fryer & Paggi, 1972 Figs. 3, 5661.
Fryer & Paggi, 1972: 255261, Figs. 113; Fryer, 1974: 204 211, Figs. 7890; Smirnov, 1976: 148, Fig. 133; 1988: 76, Fig. 35;

Type locality: Madrejуn Don Felipe also known as Laguna Ramirez a lagoon lying between the Rio Colastine and the Rio Santa Fe, between the towns of Santa Fe and Paranб [Fryer & Paggi, 1972], Argentina. Parthenogenetic female: Body high, almost globular, with slight dorsal keel. Dorsal margin arched from tip of rostrum to broadly round postero-dorsal angle, border between head and valves not expressed. Reticulation on valves and head, rows of setules arising from striae of reticulation, due to this dorsal margin squamose. Head small, with straight ventral margin and convex dorsal margin, massive supra-ocular dome and a relatively large projection for dorsal head pore elevated under


Cladocera from Pantanal
the latter, rostrum short, with frontal head pore. Valves in general ovoid, a projection on ventral margin subdivides it into two seta-fringed flanges (see Fryer & Paggi [1972]); at anterior margin of valve there is an inwardly directed hooklike projection. Postabdomen bilobed, postabdominal seta short with a relatively long distal segment. Postabdominal claws relatively large. Antenna I, II and thoracic limbs described in detail by Silva-Briano [1998] and Silva-Briano & Dumont [2001]. Size 570900 чm in our material, but up to 1100 чm according to Fryer & Paggi [1972]. Ephippial female and male unknown. COMMENTS. Silva-Briano & Dumont [2001] have serious doubts in the validity of the genus Onchobunops Fryer & Paggi, 1972, and their argument based on the similarity of all thoracic limbs in O. tuberculatus and Bunops serricaudata (Daday, 1884) is quite convincing. Recently the system of macrothricid-like anomopods (in terms of Silva-Briano [1998]) has been under reconstruction (see Dumont & SilvaBriano [1998]), and we are not sure about the necessity of immediate rejection of conventional generic names. There are problems with the species name also: previous specimens described as Macrothrix gibbera Daday, 1905 are most probably this taxon, and gibbera is a pretender to tuberculatus keeping in mind the impossibility of replacing of the major synonym in this situation, according to paragraph 23.9.1. of ICZN [2000]. Unfortunately, Dadays type specimen (DAD II/ p-467) is in a very poor condition [Smirnov, 1992], and cannot be used to clarify this difficult taxonomic situation. The situation could be resolved by an investigation of animals from the exact type locality as reported by Daday [1905], but, most probably, conservation of tuberculatus must be confirmed in particular by the International Commission on Zoological Nomenclature.

85

COMMENTS: Alona ossiani is a newly described Neotropical species from the A. affinis group, which was described based on Sars specimens from Sгo Paulo [Sinev, 1998]. We found a single specimen as well as several shell remains that could definitely be identified by the well-developed posterodorsal corner of the carapace and the setules at the posteroventral corner. The position of the head pores as well as the carapace covered with fine striae are further characteristics of Alona ossiani.

9. Karualona muelleri (Richard, 1897) Fig. 8.
COMMENTS: Dumont & Silva-Briano [2000] established a new genus Karualona, and recently the South American species Alona muelleri Richard, 1897 has been redescribed and translocated into this genus [Sinev & Hollwedel, 2003].

10. Leydigia cf. striata Birabйn, 1939
Daday, 1905: 185, Pl. 11: Fig. 19 (acanthocercoides); Birabйn, 1939: 661663, Figs. 1416; Harding, 1955: 242243, Figs. 5060 (ciliata); Smirnov, 1971: 454, 458 (Neotropical records).

COMMENTS: Harding [1955] considered this species a junior synonym of L. ciliata Gauthier, 1939, but recently Kotov et al. [2003b] demonstrated differences between these two species.

11. Oxyurella longicaudis (Birge, 1910) Figs. 7, 6271.
Daday, 1905: 178180, Pl. 11: Figs. 1213 (Euryalona tenuicaudis); Birge, 1910: 10451048, Pl. 71: Figs. 34, 7 (Odontalona); Brehm & Thomsen, 1936: 216 (Odontalona); Brehm, 1938: 100, Fig. 5 (Odontalona); Bergamin, 1941: 163, Fig. 2; Martнnez de Ferrato, 1966: 400401, Pl. 2: Figs. 34; Smirnov, 1971: 496, Figs. 627628; Elmoor-Loureiro, 1998: 37.

C. Family CHYDORIDAE Stebbing, 1902 emend. Dumont & Silva-Briano, 1998 a. Subfamily Aloninae Dybowski & Grochowski, 1894 emend. Frey, 1967 7. Alona iheringi Sars, 1901
Sars, 1901: 4951, Pl. 9: Figs. 2ac (Alona iheringi n.sp.); Sinev, 2001: 113119, Figs. 133. Not Alona davidi var. Iheringi Richard, 1897: 294296, Figs. 4243.

COMMENTS: This species of the Alona costata-group, which is closely related to A. rustica, was redescribed by Sinev [2001]. He examined Sars type material from Sгo Paulo and additional material from Boa Vista. We found only one female in the Canal do Tamengo, with external morphology that agrees with the characteristic features of this species. A. iheringi Sars, 1901 is an unacceptable taxon, a junior synonym of A. iheringi Richard, 1897, which was not mentioned by either Sars [1901] or Sinev [2001]. This taxon must be renamed, and, most probably, should be described as a new species.

8. Alona ossiani Sinev, 1998 Figs. 4, 9.
Sars, 1901: 4849, Pl. 9: Figs. 1ac (affinis); Daday, 1905: 171 (affinis); Stingelin, 1913: 621; Birabйn, 1939: 656658, Figs. 78 (affinis); Elmoor-Loureiro, 1998: 33 (Biapertura affinis); Sinev, 1998: 105110, Figs. 14.

Type locality: Not clearly indicated: Lake Charles, La., from U.S.A., and Demerara, South America, from unknown country [Birge, 1910: 1046]. Also, this author referred to Dadays [1905] description of Euryalona tenuicaudis from Paraguay as O. longicaudis, so Daday's specimens are included as members of the Birges type series according to ICZN [2000]! The type locality must be determined by means of selection of the lectotype (from materials of Birge or Daday). Parthenogenetic female: Body ovoid, elongated, compressed laterally. Dorsal margin regularly arched from tip of rostrum to practically completely smooth postero-dorsal angle, posterior margin regularly convex. Reticulation absent on head shield, while well-developed longitudinal striation present on valves. Head with relatively long rostrum, head shield somewhat elongated, with three major head pores without connection, postpore distance somewhat more than interpore distance, ocellus somewhat smaller than eye. Lateral head pores at distance similar to interpore distance from middle line of head shield. Labral keel triangular-round, large, with expressed apex, rows of setules in posterior portion of labrum, distal labral plate small. Valves ovoid, with numerous ventral marginal setae, fine setules submarginally on inner face of valve in region of postero-ventral angle and whole posterior margin. Postabdomen elongated, narrowing somewhat distally, with anal margin somewhat shorter than preanal margin, thin teeth along all postanal margin, only two distalmost teeth significantly larger than the rest, lateral fasicles of setules decreasing in size distally, and absent near largest


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Figs. 6271. Oxyurella longicaudis, parthenogenetic female from Canal do Tamengo: 62 adult in lateral view; 6364 head shield and head pores; 65 labrum; 66 setae at postero-ventral portion of valve; 6769 postabdomen, its postanal margin, and postabdominal claws; 70 antenna II; 71 thoracic limb I. Scales: 100 чm. Рис. 6271. Oxyurella longicaudis, партеногенетическая самка из Canal do Tamengo: 62 вид сбоку; 6364 головной щит и головные поры; 65 лябрум; 66 щетинки в задне-брюшной части створки; 6769 постабдомен, его постанальный край и постабдоминальный коготь; 70 антенна II; 71 нога I. Масштаб: 100 чm.

teeth, a deep embayment under base of claws. Postabdominal setae short, postabdominal claws relatively thin, curved distally, with a large basal spine located relatively far from base of claw. Antenna I not reaching tip of rostrum, with 9 aesthetascs. Antenna II relatively short, but thin, two sensory setae on coxal portion long, somewhat different in size, basal segment robust, with numerous rows of setules, branches

relatively thin. Antennal formula: setae 0-0-3/1-1-3, spines 10-1/0-0-1, all spines relatively long. Limb I with ODL bearing a single long seta, and IDL with 2 setae and a rudiment of a third seta, no gnathobase on limb I. Size 520635 чm in our material, maximal size more than that reported by Smirnov [1971]. Ephippial female: Unknown.


Cladocera from Pantanal
Male: Described poorly by Brehm [1938]. COMMENTS. This species is well-defined, and South and North American populations seem to belong to the same species. They are easily distinguished from other Neotropical species, O. ciliata Bergamin, 1939 (see Rey & Vasquez [1986]; Dimas Flores [2002]) in lacking setules on the labral keel. In contrast to O. tenuicaudis (Sars, 1862) and O. brevicaudis Michael & Frey, 1983, O. longicaudis has two especially robust teeth in distal portion of postabdomen and third seta on IDL.

87

13. Ephemeroporus hybridus (Daday, 1905) Figs. 8596.
Daday, 1905: 158159, Pl. 10: Figs. 57 (Chydorus); Stingelin, 1913: 629630, Fig. 31 (Chydorus); Martнnez de Ferrato, 1967: 329330, Pl. 1: Fig. 11 (Chydorus); Smirnov, 1971: 306, Fig. 336 (Chydorus); Green, 1972: 220 (Chydorus); Frey, 1982: 238243, Pl. 2: Figs. 18, Pl. III: Figs. 118, Pl. 4: Figs. 120; Rey & Vasquez, 1986: 152: Pl. 7: Figs. 112; Smirnov, 1996: 155156, Figs. 643 648; Elmoor-Loureiro, 1998: 29.

b. Subfamily Chydorinae Stebbing, 1902 12. Alonella dadayi (Birge, 1910) Figs. 7284.
Daday, 1905: 168169, Pl. 10: Figs. 1823 (Leptorhynchus dentifer); Stingelin, 1909: 644, Fig. 2 (Leptorhynchus dentifer); Birge, 1910: 1052; Brehm & Thomsen, 1936: 217, Fig. 9; Martнnez de Ferrato, 1966: 398399, Pl. 1: Figs. 57; Smirnov, 1971: 277, Fig. 290 (Disparalona); Green, 1972: 221; Paggi, 1995: Fig. 127 (Phryxura); Smirnov, 1996: 8890, Figs. 328331; Elmoor-Loureiro, 1998: 29 (Disparalona); Dimas Flores, 2002: 77, Pl. 1: Figs. AF.

Type locality: Not reported accurately in the first description, only marked as Louisiana, Tex. in a table on p. 1053. Also Birge [1910] referred to Leptorhynchus dentifer Daday, 1905, so Dadays material can be regarded a part of the Birges type series according to ICZN [2000]! It is necessary to select a lectotype of this species also. Parthenogenetic female: Body ovoid, moderately elongated, compressed laterally. Dorsal margin regularly arched from tip of rostrum to distinct postero-dorsal angle, posterior margin truncated, postero-ventral angle rounded, with series of small denticles, setules between them. Whole shell richly sculptured with longitudinal striation in dorsal portion and head shield, and polygonal reticulation in posterior portion of valves, parallel strokes within reticulation cells. Head large, with remarkably long, somewhat curved backwards rostrum, ocellus smaller than eye. Head pores on special narrow, nonreticulated plate, each of two major head pores with thin ring, postpore distance only 1/4 interpore distance, two minute pores on mid-line, somewhat closer to anterior major pore. Labral keel large, more or less triangular-round, with wavy anterior margin and apex directed posteriorly, distal labral plate relatively large. Postabdomen robust, high, narrowing distally, preanal margin much longer than anal margin, preanal angle inflated, obtuse or right angled, a series of especially long setules in this region; anal margin significantly concave, with small, numerous teeth. Postanal margin shorter than preanal margin, with 68 teeth. Postabdominal seta shorter than postabdomen, its distal segment shorter than basal one, with relatively long setules; postabdominal claws relatively short and thick, with robust basal spine at a distance from claw base, and a small additional spine closer to claw base. Antenna I thick, with fine, short sensory seta and 9 aesthetascs of different size. Antenna II relaticvely short, but elongated, antennal formula: setae 0-0-3/1-1-3, spines 1-0-1/0-0-1. Each swimming seta with distal segment armed with setules, and chitinous insertions near joint with basal segment. ODL I with a single naked seta, IDL with 3 setae, smallest naked. Size 240270 чm in our material, up to 450 чm according to Smirnov [1996]. Ephippial female: unknown. Male: Smirnov [1996] only gave the size as 200 чm. COMMENTS. This is a very typical neotropical species of Alonella, remarkably different from others (see Smirnov [1996]).

Type locality: there is only a note Paraguay on labels of Dadays types in HNHM. Also, in his text, the author did not select the locality from the series of water bodies investigated, so the exact type locality is not clear [Smirnov, 1996]. Parthenogenetic female: Body globular, high and thick, dorsal margin regularly arched from tip of rostrum to a rounded, but visible, postero-dorsal angle, posterior margin straight or concave. Postero-ventral angle widely rounded, with single denticle, but in many females from Pantanal this denticle significantly or completely reduced, and with specialized short setae dorsally to the tooth. Head with welldeveloped rostrum directed downwards, ocellus somewhat smaller than eye. No head pores in adult state. Labral keel projected ventrally, with well-developed apex and single tooth at anterior margin (in a single female this tooth was greatly reduced). Valves ovoid, with polygonal reticulation, numerous short lines within cells of reticulation, setae in anterior portion of ventral edge marginal, in the middle absent, in posterior portion submarginal, densely setulated. Postabdomen elongated, narrowing somewhat distally, dorso-distal angle rounded, distal end truncated. Preanal margin long, preanal angle prominent, postanal margin with large teeth, 3 proximal anal teeth especially long. Postabdominal seta shorter than postabdomen, postabdominal claw regularly curved, with a long basal spine and a minute additional basal spine. Antennal formula: setae 0-0-3/ 0-1-3, spines 1-0-1/0-01, all spines very short. Limb I with IDL bearing a single seta, IDL with 3 seta of different size, gnathobase I with a single seta and setulated projection. Size 220270 чm in our material, up to 320 чm according to Rey & Vasquez [1986]. Ephippial female: See Frey [1982]. Male: still unknown [Smirnov, 1996] COMMENTS. Our E. hybridus was collected close to the presumed type locality. Our main conclusion is that it is necessary to be very careful with presence-absence of denticles at postero-ventral angle of valves as a key character for the discrimination of species within the genusEphemeroporus, because in Pantanal populations these were reduced in many specimens. A similar situation was observed with another diagnostic feature, a tooth on labrum: in one specimen from Pantanal the tooth on the labrum was strongly reduced in size. In addition, Frey [1982] noted that there are several species in the world, not yet described, allied to E. hybridus.

14. Pseudochydorus globosus (Baird, 1843) Figs. 97108.
Bergamin, 1940: 99, Fig. 18 (Chydorus); Martнnez de Ferrato, 1967: 328329, Pl. 1: Figs. 67; Paggi, 1995: Figs. 104106; Elmoor-Loureiro, 1998: 31. Not Rey, 1993: 259, Figs. 2627.

Type locality: Ditch near Richmond; pond near Isleworth, United Kingdom, Europe. Parthenogenetic female: Body subglobular to globular, thick, dorsal margin regularly arched from tip of rostrum to posterior margin, sometimes there is a slight prominence


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Werner Hollwedel, Alexey A. Kotov & Gerd-Oltmann Brandorff

Figs. 7284. Alonella dadayi, parthenogenetic female from Baia 38: 72 adult in lateral view; 73 head pores; 7475 labrum; 7677 postero-ventral portion of valve in external view; 78 the same in internal view; 7980 postabdomen; 81 antenna I; 8283 antenna II and joint part of its swimming seta; 84 distal portion of thoracic limb I. Scales: 100 чm. Рис. 7284. Alonella dadayi, партеногенетическая самка из Baia 38: 72 вид сбоку; 73 головные поры; 7475 лябрум; 7677 задне-брюшная часть створку снаружи; 78 изнутри; 7980 постабдомен; 81 антенна I; 8283 антенна II и сочленение дистального и базального члеников плавательной щетинки; 84 дистальная часть ноги I. Масштаб: 100 чm.


Cladocera from Pantanal

89

Figs. 8596. Ephemeroporus hybridus, parthenogenetic female from Baia 38: 85 adult in lateral view; 8687 labrum; 88 89 valve and its sculpture; 9091 setae at middle and posterior portions of valve ventral margin; 9294 armature of posteroventral angle in 3 valves; 95 postabdomen; 96 thoracic limb I. Scales: 100 чm. Рис. 8596. Ephemeroporus hybridus, партеногенетическая самка из Baia 38: 85 вид сбоку; 8687 лябрум; 8889 створка и ее скульптура; 9091 щетинки в середине и задней части брюшного края; 9294 вооружение задне-брюшного угла трех разных створок; 95 постабдомен; 96 нога I. Масштаб: 100 чm.

under compound eye, postero-dorsal angle smooth. Reticulation not visible. Head with well-developed rostrum, two major head pores in mid-line of head shield, interpore distance greater than postpore one, small pores in mid-line also, between two major pores. Labral keel reduced. Valves ovoid, with setae submarginally on inner face along entire

ventral margin. Postabdomen elongated, with parallel dorsal and ventral margins, preanal margin longer than anal margin, preanal angle somewhat prominent, postanal margin with small teeth, distal angle rounded. Basis of postabdominal claws remarkably prominent distally, postabdominal claw relatively long and thin, with a single basal spine immediately


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Werner Hollwedel, Alexey A. Kotov & Gerd-Oltmann Brandorff

Figs. 97108. Pseudochydorus globosus, parthenogenetic female from Canal do Tamengo: 97 adult in lateral view; 98 head pores; 99102 armature of valve, and its antero-ventral, ventral and postero-ventral portions; 103 distal portion of postabdomen; 104105 antenna II and joint portion of its swimming seta; 106 maxilla I; 107108 thoracic limb I and its distal portion. Scales: 100 чm. Рис. 97108. Pseudochydorus globosus, партеногенетическая самка из Canal do Tamengo: 97 вид сбоку; 98 головные поры; 99102 вооружение створки, ее передне-брюшногй, брюшной и задне-брюшной части; 103 дистальная часть постабдомена; 104105 антенна II и сочленение дистального и базального члеников ее плавательной щетинки; 106 максиллаI; 107108 нога I и ее дистальная часть. Масштаб: 100 чm.


Cladocera from Pantanal
at its base, and spaced denticles along its dorsal margin. Antenna I short, robust, with 9 aesthetascs of different size. Antenna II relatively small, basal segment and segments of branches robust. Antennal formula: setae 0-0-3/1-1-3 (seta at basal endopod segment very short), spines 0-0-1/0-0-1, apical spines minute. Distal segments of swimming setae setulated, with chitinous insertions near joints with basal segments. No filter setae on any limbs or maxillae I. Limb I with ODL bearing a long and a short seta, IDL with three setae, shortest seta hook-like, and a bunch of robust setules. Other limbs not different from those described by Smirnov [1971]. Size 400450 чm in our material. Ephippial female, male: Unknown from the Neotropics. COMMENTS. The size of the labral keel, which is used in all keys for the discrimination of Chydorus and Pseudochydorus, is in fact not a major difference between these genera; structure of limbs is a far more important trait; in Pantanal females all limbs were clearly non-filtering. We did not find any significant differences in the structure of thoracic limbs in Pantanal and European females, but our ideas on the latter were based on pictures of Fryer [1968] and Smirnov [1971], published around 30 years ago, that do not comply with recent standards of anomopod descriptions. The status of South American populations should be checked after a redescription of European P. globosus s. str. In contrast, Rey [1993: 259, Figs 2627] most probably describedChydorus, not Pseudochydorus, from Lake Titicaca: the labral keel in her animal was small but existing, with expressed apex, and the postabdomen was not of the Pseudochydorus-type.

91

surveys only examined a limited number of water bodies about 10 or even fewer. Thus the majority of Brazil is waiting for thorough investigation of its cladoceran fauna. Many previously established species, European as well as tropical, must be redescribed according to the new standards [Korovchinsky, 1996]. Not only rare but many very common Brazilian, other South American, and Australian species of the Anomopoda are described very poorly, and this has resulted in continual misidentifications.
ACKNOWLEDGEMENTS: We are very grateful to Dr H. J. G. Dartnall and Prof. R. J. Shiel for editing an earlier draft, to Prof. N. N. Smirnov for valuable consultations, to Dr I. B. Mertsalov for help in preparation of material for SEM, Mr. V. N. Antropov for technical assistance in the work with SEM, Mr. H. Bahena-Basave for the artwork in photographs, and Mбrcia Divina de Oliveira and Dйbora Fernandes Calheiros for the access to the samples from the Rio Paraguay and Baнa de Castelo. GOB thanks the Empresa Brasileira de Pesquisa Agropecuбria (EMBRAPA), Centro de Pesquisa Agropecuбria do Pantanal (CPAP) for kind support of his travel to Brazil. AAK started preparation of illustrations during his stay in Mexico, and he thanks the Consejo Nacional de Ciencia y Tecnologia (CONACYT) for supporting his stay in Mexico, through the Catedras Patrimoniales program, and to Prof M. Elнas-Gutiйrrez for logistic help during the stay.

Discussion
The Pantanal fauna is quite typical of Brazil and Neotropical regions. The similarity of our species list with ones already published for these regions is quite obvious, especially bearing in mind that some differences can be explained by the changing taxonomic status of some species. Although no new species are presented in this paper, we have indicated many areas of confusion with European forms (see above) that require clarification, but they are not endemics of the Pantanal. Large South American rivers such as the Rio Paraguay flow through thousands of kilometres, and are important arteries for dispersion of freshwater species; as are the myriads of aquatic birds associated with the river banks that participate in dispersion of anomopod ephippia. The definitive list of Brazilian Cladocera is far from complete, with new species being described all the time [Smirnov & Santos-Silva, 1995; Sinev & Hollwedel, 2002]. Brazilian investigators have published many lists of Cladocera from different regions, including the Pantanal [Lima et al., 1996, 1998] but detailed investigations including descriptions of some species have only been made for a few regions. These regions include the vicinity of Sгo Paulo, Itatiba and Ipiranga (Sгo Paulo State) [Sars, 1901], the city of Sгo Paulo [Bergamin, 1940, 1941], the region of Rio Nhamundб (border of Parб and Amazonas) [Brandorff et al., 1982], Lagoa dos Patos (Rio Grande do Sul) [Montъ & Gloeden, 1986], the region of Boa Vista (Roraima) [Smirnov & SantosSilva, 1995], and Lenзуis Maranhenses dune field (Maranhгo) [Van Damme, in preparation]. Most of these

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